植物科学最前線

6: 23 ( 2015)

 $&   'ź¯ ŦøĊƅ ÝŚř ĮøƔÄúƧưıƕðǸƂƫúƧưXzZ|230-0051 ƪöċƣůſčȱǦÅŘēƙ 2-7-11
How do wounded plants repair or regenerate?
Key words: regenerative capacity, repair, wounding
Momoko Ikeuchi, Akira Iwase, Keiko Sugimoto
RIKEN Center for Sustainable Resource Science
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2. *+ !)/
2-1. #(.*+' ť/¨Ƴ-0, ȣųĤȲQuiescent Center, QCȳ<2
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ä 1. ťƳµǢƾǈ/ŮĖäťƳ=130 µm )¶Ȟ(:ƺ 80%/ȰÁƒ)°Ɩ)?Sena et al., 2009 <>ǬÑC
ġ(ǿȂBerg et al., 1995) /Ȣ-0ƻǐƹǵ0ƇȜ) >,Oy|ew,ǉīëģ§ȒƭCÒ%( ?:/*ǌ =@?/ǀǳ0, Ēƻǐb\[ªCÖ8ťƳȩèC¶Ȟ?þȯƹ- (:ŁĶ@( ?ťƳ¶Ȟğ, ƾǈ/°ǆįCƿ
M. Ikeuchi, A. Iwase & K. Sugimoto-1
BSJ- Revi ew 6: 23 ( 2015)

植物科学最前線

Quiescent centre

Columella

Xylem

St

Vasculature

70 µm

6: 24 ( 2015)

Stem cells(initials)

Frequency of regeneration–4 –2 0 +2 +4b(ńŌ¯-ťƳȩè°Ɩ@?,/°Ɩ- (Ēƻǐb\[/ŰǑ-ĥȨ,ǿ²âø
1.0

Quiescent centre) ? PLETHORA1(PLT1), PLT2, SCARECLOW (SCR) 0ĥǥ,, ŋù/ǉīëȲ =
0.8

0.6
K|MUzƄĕÃȗȳ-ê'(ťƳµǢƾǈ/}ǗƝ,ƻǐviyOunzO( QC ;Ē
0.4
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$,7-,
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est
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roo
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pro
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enr
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or
ind

Distance from tip (µm)

c

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0 d.p.c.

1 d.p.c.

2 d.p.c.

3 d.p.c.

4 d.p.c.

CYCB1;1::GFP
3 d.p.c. et al., 2009
4 d.p.c.
0 d.p.c.
1 d.p.c.
2 d.p.c.
Uncutä 2.dUyGcaWaťƳ/°ƖCYCB1;1GFP)µǢƻǐm|N@(?Sena<>ǬÑCġ(ǿȂ

PET1112-2. ,0.*+'
0 d.p.c.
1 d.p.c.
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2 d.p.c.
 Ǜ/¨Ƴ?ǛȧµǢƾǈ)0, ƇȝįȚCǟ ŹǃĪ*3 d.p.c.

4 d.p.c.

ň",àüCƖ7´ěĭěįǑCµǢƾǈ¯/ƚ,?ȩèµĴ( ?`|o/ȧƳȖ-ǉ? central zone (CZ) ĒƻǐCÖ8ȩè)?/-Ą, ȆǇȖ-ǉ? peripheral
pWOX5::GFP
zone (PZ) 0 ƖàüȲǝ;ǘ,+ȳ/ÉêCƖ7´ĝ½CĴ
13,14

for3ȳ
root
gravitropism CZ. By
one day post
Lugol staining confirmedȲäǛȧµǢƾǈ/Ľ£?*,
PZcut,
=}&,

germination quickly regene
de novo starch accumulation above the cut site (Fig. 2a andvergent pattern of longitu
Ǥń/µǢƾǈ°Ɩ?°Ɩ-¨Ʋ$, ĒƻǐŰǑCŁ
ä two
3. ǛȧµǢƾǈ/ŮĖä
Supplementary Fig. 4). More intense staining was observeddays
Supplementary Fig. 5a). M?post
organizing
center(OC))ƜƓ?
WUSCHEL(WUS)/Ɯ
Reinhardt of
et al.,the
2003
<>ǬÑ
cut (Fig. 2a and Supplementary Fig. 4). To test for recovery
regenerating
double mucolumella function, we subjected regeneratingroots to aetstandard
graviwasre-established(Fig. 2cƓȖ°ǆį@?*Ǩă@(? (ä 4; ReinhardtCġ(łñ
tropism assay by reorienting them perpendicularly to the gravity vector
functional columella cellsal., 2003)ǛȧµǢƾǈ/°Ɩ0, Ēƻǐb\[õB@(:Øand scoring the response over time. All wild-type roots showed a clear
Similarly, scarecrow (scr) mu
ã/µǢȩè/ƻǐviyOunzOCƿ(Ēƻǐb\[C°Ɩ?*Ɔ-(ťƳ/gravitropic response within 12 h. Although cut roots did not respond tofunctionthrough a pathway
gravity in the?*Ǫ
first 12 h?@6)/Ƨư)0,
after excision when cut at
130 mm, 13.8% of the17), were also able to restor°Ɩ*«ȋ(þȯśņ*(_m_,+ǛȧŶȁƝôroots
exhibiteda clear
response
1 day
cut,
55.4%
gravitropism cutĿ; ŧƌ Ƙgravitropic=@(" " 9 ,atȕ úpostƝ,ƥǦǻ( ?  , (Fig. 2b–d, f an
at 2 days post cut and 89.2% at 3 days post cut (n 5 65, for all time
PLT2 are expressed early in
WUS-CLAVATA(CLV) /ÍĦĵŃq^w-<%(°Ɩ/fZ|z<ǲō)?
Ȳä 5; Fujita
points). However,the quiescent-centre-specific marker WUSCHEL
mentary
Fig. 7). However,
RELATED HOMEOBOX 5 (WOX5) was either ectopically expressedmutant and wild-type roo
in the endodermal file or, at times, expressed in differentiated columelladependent genes were induccells at one day post cut (Fig. 2a and Supplementary Fig. 4). Thus, as
ing double mutants (Suppearly as one day after complete columella excision, a new set of cells
percentage of plt1plt2 and s
expressed columella markers and performed columella-specific functype roots (Fig. 2c), whichtions while the morphology of the stem cell niche had not yet recovered.
effect of both mutants in r
zone15,17—the pool of cells
Given the early re-establishment of a differentiated cell type, we
results show that stem cell ntested the requirement for functional stem cells by using mutants in
tip regeneration and imply twhich post-embryonic root growth ceases due to the failure to mainä 4. tain_m_Ǜȧ-?
LeWUS/ƜƓfZ|zOC/ƻǐõB@?*(B),
1
Ōğ-0ØȆ)Ę
for cell specification and pa
the stem cell niche. The PLETHORA (PLT) gene family has beenwiththe double
Several results suggest th
shown tocritical for root formation15, (D)ƜƓ÷6>(C),be2 Ōğ-0ƜƓȖ/°ŬįǺ?
Reinhardt
et al., mutant
2003 <>ǬÑCġ(ǿȂ
plt1plt2 showing differentiation of stem cells at three days post gerthe root stem cell niche and
under
our
conditions
(Fig.
2b,
note
the
lack
of
mination5, as verified M.
tration gradient18,19, may be
Ikeuchi, A. Iwase & K. Sugimoto-2
stem cell layer between the quiescent centre and the starch-stained
coordinates organogenesis2
earliest
stagescolumella). The uncut double mutant root has abnormal tip and stem BSJRevi ew
6: 24when( 2015)we blo
using N-1-naphthylphthalacell niche morphology but normal gravitropism and convergent lon5

植物科学最前線

6: 25 ( 2015)

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BSJ- Revi ew 6: 25 ( 2015)

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BSJ- Revi ew 6: 26 ( 2015)

植物科学最前線

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BSJ- Revi ew 6: 27 ( 2015)

植物科学最前線

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ėƘŅƐ
Aida, M., Beis, D., Heidstra, R., Willemsen, V., Blilou, I., Galinha C., Nussaume L, Noh Y-S, Amasino R,
and Scheres B (2004). The PLETHORA genes mediate patterning of the Arabidopsis root stem cellniche. Cell 119: 109-120.
Birnbaum, K.D. and Sánchez Alvarado, A. (2008). Slicing across kingdoms: regeneration in plants andanimals. Cell 132: 697–710.
Fujita, H., Toyokura, K., Okada, K., and Kawaguchi, M. (2011). Reaction-diffusion pattern in shoot apicalmeristem of plants. PLoS ONE 6: e18243.
Ikeuchi, M., Igarashi, H., Okada, K., and Tsukaya, H. (2014). Acropetal leaflet initiation of Eschscholzia
californica is achieved by constant spacing of leaflets and differential growth of leaf. Planta
240:125-135.
Iwase, A., Mitsuda, N., Koyama, T., Hiratsu, K., Kojima, M., Arai, T., Inoue, Y., Seki, M., Sakakibara, H.,
Sugimoto, K., and Ohme-Takagi, M. (2011). The AP2/ERF transcription factor WIND1 controls cell
dedifferentiation in Arabidopsis. Curr. Biol. 21: 508–514.
Kuchen, E.E., Fox, S., de Reuille, P.B., Kennaway, R., Bensmihen, S., Avondo, J., Calder, G.M., Southam,P., Robinson, S., Bangham, A., and Coen, E. (2012). Generation of leaf shape through early patterns ofgrowth and tissue polarity. Science 335: 1092–1096.
Reinhardt, D., Frenz, M., Mandel, T., and Kuhlemeier, C. (2003). Microsurgical and laser ablation analysis
of interactions between the zones and layers of the tomato shoot apical meristem. Development 130:4073–4083.
Sena, G., Wang, X., Liu, H.-Y., Hofhuis, H., and Birnbaum, K.D. (2009). Organ regeneration does not
require a functional stem cell niche in plants. Nature 457: 1150–1153.

M. Ikeuchi, A. Iwase & K. Sugimoto-7
BSJ- Revi ew 6: 29 ( 2015)

植物科学最前線

6: 30 ( 2015)

Sena, G and Birnbaum K.D. (2010) Built to rebuild: in search of organizing principles in plant regeneration
Curr. Opin. Genet. Dev. 20: 460–465.
van den Berg, C., Willemsen, V., Hage, W., Weisbeek, P., and Scheres, B. (1995). Cell fate in the
Arabidopsis root meristem determined by directional signalling. Nature 378: 62–65.

M. Ikeuchi, A. Iwase & K. Sugimoto-8
BSJ- Revi ew 6: 30 ( 2015)